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|Location||Cortex esp. Layers III and V|
|Function||excitatory projection neuron|
|Gray's||subject #183 722|
Pyramidal neurons (pyramidal cells) are a type of neuron found in areas of the brain including cerebral cortex, the hippocampus, and in the amygdala. Pyramidal neurons are the primary excitation units of the mammalian prefrontal cortex and the corticospinal tract. Pyramidal neurons were first discovered and studied by Santiago Ramón y Cajal. Since then, studies on pyramidal neurons have focused on topics ranging from neuroplasticity to cognition.
One of the main structural features of the pyramidal neuron is the triangular shaped soma, or cell body, after which the neuron is named. Other key structural features of the pyramidal cell are a single axon, a large apical dendrite, multiple basal dendrites, and the presence of dendritic spines. 
The apical dendrites arise from the apex of the pyramidal cell's soma. The apical dendrite is a single long thick dendrite that branches several times as distance from the soma increases. 
The basal dendrites arise from the base of the pyramidal cell's soma. The basal dendritic tree consists of three to five primary dendrites. As distance increases from the soma, the basal dendrites branch profusely. 
In the cerebral cortex, the size of the dendritic trees of pyramidal cells have been shown to differ systematically among different cortical areas. They become increasingly larger, more branched and spiny with progression through visual areas, for example. Similar trends have been demonstrated in somatosensory and motor cortex. Those in cingulate cortex are even larger and more branched. Pyramidal cells in the prefrontal cortex of man are the most spiny of all pyramidal cells. Moreover comparative studies reveal that pyramidal cells in man are more spiny than those in the chimpanzee, old world monkeys and new world monkeys. As each spine receives at least one excitatory input, the implication is that pyramidal cells in the prefrontal cortex of man are connected to more cells, and embedded in more complex circuits, than those in other cortical areas and species.
Dendritic spines receive most of the excitatory impulses (EPSPs) that enter a pyramidal cell. Dendritic spines were first noted by Ramón y Cajal in 1888 by using Golgi's method. Ramón y Cajal was also the first person to propose a physiological role of dendritic spines: increase the receptive surface area of the neuron. The greater the pyramidal cell's surface area, the greater the neuron's ability to process and integrate large amounts of information. Dendritic spines are absent on the soma, and the number of spines increases away from it. The typical apical dendrite in a rat has at least 3000 dendritic spines. The average human apical dendrite is approximately twice the length of a rat's, so the number of dendritic spines present on a human apical dendrite could be as high as 6000. 
Growth and developmentEdit
Pyramidal specification occurs during early development of the cerebrum. Progenitor cells are committed to the neuronal lineage in the subcortical proliferative ventricular zone (VZ) and the subventricular zone (SVZ). Immature pyramidal cells undergo migration to occupy the cortical plate, where they further diversify. Endocannabinoids (eCBs) are one class of molecules that have been shown to direct pyramidal cell development and axonal pathfinding. Growth factors such as Ctip2 and Sox5 have been shown to contribute to the direction in which pyramidal neurons direct their axons.
Early postnatal developmentEdit
Pyramidal cells in rats have been shown to undergo many rapid changes during early postnatal life. Between postnatal days 3 and 21, pyramidal cells have been shown to double in the size of the soma, increase in length of the apical dendrite by five fold, and increase in basal dendrite length by thirteen fold. Other changes include the lowering of the membrane’s resting potential, reduction of membrane resistance, and in increase in the peak values of action potentials.
Early postnatal development Edit
Recent studies on the development of pyramidal cells in primates reveal that at birth the dendritic trees of pyramidal cells in different cortical areas already differ in their size, number of branches and number of spines. The pyramidal cells in the different cortical areas then grow according to specific developmental profiles, resulting in the specialization observed in different cortical areas in the mature brain.
Like dendrites in most other neurons, the dendrites are generally the input areas of the neuron, while the axon is the neuron’s output. Both axons and dendrites are highly branched. The large amount of branching allows the neuron to receive and send signals to many different neurons.
Pyramidal neurons, like other neurons, have numerous voltage-gated ion channels. In pyramidal cells, there is an abundance of Na+, Ca2+, and K+ channels in the dendrites, and some channels in the soma. Ion channels within pyramidal cell dendrites have different properties from the same ion channel type within the pyramidal cell soma. Voltage-gated Ca2+ channels in pyramidal cell dendrites are activated by subthreshold EPSPs and by back-propagating action potentials. The extent of back-propagation of action potentials within pyramidal dendrites depends upon the K+ channels. K+ channels in pyramidal cell dendrites provide a mechanism for controlling the amplitude of action potentials.
The ability of pyramidal neurons to integrate information depends on the number and distribution of the synaptic inputs they receive. A single pyramidal cell receives about 30,000 excitatory inputs and 1700 inhibitory (IPSPs) inputs. Excitatory (EPSPs) inputs terminate exclusively on the dendritic spines, while inhibitory (IPSPs) inputs terminate on dendritic shafts, the soma, and even the axon. Pyramidal neurons use glutamate as their excitatory neurotransmitter, and GABA as their inhibitory neurotransmitter.
Firing Classification of Pyramidal NeuronsEdit
Pyramidal neurons have been classified into different subclasses based upon their firing responses to 400-1000 millisecond current pulses. These classification are RSad, RSna, and IB neurons.
RSad Pyramidal NeuronsEdit
RSad pyramidal neurons, or adapting regular spiking neurons, fire with individual action potentials (APs), which are followed by a hyperpolarizing afterpotential. The afterpotential increases in duration which creates spike frequency adaptation (SFA) in the neuron. 
RSna Pyramidal NeuronsEdit
RSna pyramidal neurons, or non-adapting regular spiking neurons, fire a train of action potentials after a pulse. These neurons fail to show any signs of adaptation.
IB Pyramidal NeuronsEdit
Pyramidal neurons are the primary neural cell type in the corticospinal tract. Normal motor control depends on the development of connections between the axons in the corticospinal tract and the spinal cord. Pyramidal cell axons follow cues such as growth factors to make specific connections. With proper connections, pyramidal cells take part in the circuitry responsible for vision guided motor function.
Pyramidal neurons in the prefrontal cortex are implicated in cognitive ability. In mammals, the complexity of pyramidal cells increases from posterior to anterior brain regions. The degree of complexity of pyramidal neurons is likely linked to the cognitive capabilities of different anthropoid species. Because the prefrontal cortex receives inputs from areas of the brain that are involved in processing all the sensory modalities, pyramidal cells within the prefrontal cortex may process many different types of inputs. Pyramidal cells may play a critical role in complex object recognition within the visual processing areas of the cortex.
- ↑ 1.0 1.1 Elston GN (November 2003). Cortex, cognition and the cell: new insights into the pyramidal neuron and prefrontal function. Cereb. Cortex 13 (11): 1124–38.
- ↑ 2.0 2.1 García-López P, García-Marín V, Freire M (November 2006). Three-dimensional reconstruction and quantitative study of a pyramidal cell of a Cajal histological preparation. J. Neurosci. 26 (44): 11249–52.
- ↑ 3.0 3.1 3.2 3.3 Megías M, Emri Z, Freund TF, Gulyás AI (2001). Total number and distribution of inhibitory and excitatory synapses on hippocampal CA1 pyramidal cells. Neuroscience 102 (3): 527–40.
- ↑ Elston GN (2003) Cortex, cognition and the cell: new insights into the pyramidal neuron and prefrontal function. Cerebral Cortex 13: 1124-1138 Jacobs B, Scheibel AB (2002) Regional dendritic variation in primate cortical pyramidal cells. In: Cortical Areas: Unity And Diversity. (Schüz A, Miller R, eds.), pp 111-131. London: Taylor and Francis.
- ↑ Elston GN, Tweedale R, Rosa MGP (1999) Cortical integration in the visual system of the macaque monkey: large scale morphological differences of pyramidal neurones in the occipital, parietal and temporal lobes. Proc R Soc Lond Ser B 266: 1367-1374
- ↑ Elston G.N., Rockland K. (2002) The pyramidal cell in somatosensory and motor cortex of the macaque monkey: phenotypic variation. Cerebral Cortex 12: 1071-1078.
- ↑ Elston G.N.,Benavides-Piccione R. and DeFelipe J. (2005) The pyramidal cell in the cingulate gyrus of the macaque monkey with comparative notes on inferotemporal and primary visual cortex. Cerebral Cortex 15:64–73
- ↑ Elston, G.N. (2003) Cortex, cognition and the cell: new insights into the pyramidal neuron and prefrontal function. Cerebral Cortex 13: 1124-1138.
- ↑ Elston G. N., Benavides-Piccione R. and DeFelipe J. (2001) The pyramidal cell in cognition: a comparative study in human and monkey. Journal of Neuroscience. 21:RC163(1-5).
- ↑ Arellano JI, Espinosa A, Fairen A, Yuste R, DeFelipe J (2007) Non-synaptic dendritic spines in neocortex. Neuroscience 145: 464-469
- ↑ Elston G.N. (2007) Evolution of the pyramidal cell in primates. In: Evolution of Nervous Systems, Volume IV. Eds Kaas JH, Preuss TD. Academic Press. Oxford. 191-242.
- ↑ Laberge D, Kasevich R (November 2007). The apical dendrite theory of consciousness. Neural Netw 20 (9): 1004–20.
- ↑ Mulder J, Aguado T, Keimpema E, et al. (June 2008). Endocannabinoid signaling controls pyramidal cell specification and long-range axon patterning. Proc. Natl. Acad. Sci. U.S.A. 105 (25): 8760–5.
- ↑ Fishell G, Hanashima C (February 2008). Pyramidal neurons grow up and change their mind. Neuron 57 (3): 333–8.
- ↑ Zhang ZW (March 2004). Maturation of layer V pyramidal neurons in the rat prefrontal cortex: intrinsic properties and synaptic function. J. Neurophysiol. 91 (3): 1171–82.
- ↑ Elston G.N., Oga T., Fujita I. (2009) Spinogenesis and pruning scales among functional hierarchies. J Neurosci 29:3271-3275
- ↑ Elston G.N., Oga T., Okamoto T., Fujita I. (2009) Spinogenesis and pruning from early visual onset to adulthood: An intracellular injection study of layer III pyramidal cells in the ventral visual cortical pathway of the macaque monkey. Cerebral Cortex 20: 1398-1408.
- ↑ Elston G.N., Okamoto T., Oga T., Dornan D., Fujita I. (2010) Spinogenesis and pruning in the primary auditory cortex of the macaque monkey (Macaca fascicularis): An intracellular injection study of layer III pyramidal cells. Brain Research 1316: 35-42.
- ↑ Elston, G. N., Oga T., Okamoto T. and Fujita I. (2011). Spinogenesis and pruning in the anterior ventral inferotemporal cortex of the macaque monkey: An intracellular injection study of layer III pyramidal cells Front Neuroanat. 5:42.
- ↑ Elston GN, Fujita I. (2014) Pyramidal cell development: postnatal spinogenesis, dendritic growth, axon growth, and electrophysiology. Front Neuroanat. 8:78. doi: 10.3389/fnana.2014.00078
- ↑ Magee J, Hoffman D, Colbert C, Johnston D (1998). Electrical and calcium signaling in dendrites of hippocampal pyramidal neurons. Annu. Rev. Physiol. 60: 327–46.
- ↑ 22.0 22.1 22.2 Franceschetti S, Sancini G, Panzica F, Radici C, Avanzini G (April 1998). Postnatal differentiation of firing properties and morphological characteristics in layer V pyramidal neurons of the sensorimotor cortex. Neuroscience 83 (4): 1013–24.
- ↑ Salimi I, Friel KM, Martin JH (July 2008). Pyramidal tract stimulation restores normal corticospinal tract connections and visuomotor skill after early postnatal motor cortex activity blockade. J. Neurosci. 28 (29): 7426–34.
- Pyramidal cell - Cell Centered Database
- Diagram (as part of slideshow)
- NIF Search - Pyramidal Cell via the Neuroscience Information Framework
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